INTRODUCTION to ARTIODACTYLA (Even-Toed Ungulates)
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INTRODUCTION TO ARTIODACTYLA (even-toed ungulates) Ungulates (meaning roughly "being hoofed" or "hoofed animal") are mammals, which are herbivorous, terrestrial and relatively large in size. Most ungulates use the tips of their toes, to sustain their whole body weight while moving. A hoof is really just a modified toenail. Unlike claws and nails, hooves are the principal point of contact between the legs and the ground. The ungulates consists of two orders: Perissodactyla (odd-toed ungulates) and Artiodactyla (even- toed ungulates). Artiodactyl’s weight is borne about equally by the third and fourth toes. Artiodactyla comes from the Greek meaning even (artio) toed (dactyl). During their evolution ungulates developed hooves instead of claws. The two orders that exist today diverged from a common hoofed ancestor 60 million years ago during the Eocene period. Evolving along a different course the artiodactyls surpassed the perissodactyls in number and continues to thrive, while the perissodactyls are slowly becoming extinct. Europe was probably the center of evolution of this order. The earliest even-toed ungulates evolved in the Eocene, about 54 million years ago. These early artiodactyls had a full complement of teeth (a total of 44), four distinct digits on each foot, separate foot bones, no frontal appendage, and a simple, non-ruminating stomach. The even-toed ungulates have come into their prime only in recent times. With 77 genera, they represent the largest number of present-day ungulates and the largest number of large mammals. Most of the world’s prey animals and nearly all human-related important domesticated species come from the artiodactyla order of mammals, including pigs, camels, llamas, reindeer, yak, cattle, water-buffalo, goats and sheep. From prehistory times they have provided emerging Homo sapiens with food, drink, clothing and transport, enabling the development from primitive hunter- gather to agrarian societies. TAXONOMY The Artiodactyla Order is currently composed of approximately 160 species divided into 9 families and 77 genera. They are native to all land areas of the world, except the West Indies, Australia, New Zealand and most oceanic islands. These figures will vary depending upon which zoologists that is doing the classification. All are agreed that are 9 families. What follows is based upon Grzimek (1991). Suidae (pigs) Tayassuidae (peccary, javalina) Hippopotamidae (hippopotamus) Camelidae (camels, llamas) Tragulidae (chevrotain, mouse deer) Gifaffidae (giraffe, okapi) Cervidae (deer) Bovidae (antelope, bison, cattle, sheep, goats) Antelocapridae (pronghorn) Physical Adaptations The definitive characteristic of the even-toed ungulates is the main axis of the foot passes between the third and forth digits, which have hooves. Most members of this order walk on the tips of their toes with the weight evenly supported on the third and forth digit while the second and fifth are either reduced or lost. The metacarpus and metatarsus of the two main digits are usually merged to form the ‘cannon bone’. 1 1 1 2 2 2 1. Metacarpal (Cannon) bone 2. Phalanges A B C D E F Pigs Deer Camels Deer Camel Hippopotamus In pigs (A) the 2nd and 5th digits are still well developed and the metacarpal bones are not fused. In deer (B) the 2nd and 5th digits are vestigial and the metacarpals have fused into the cannon bone. Camels (C) no longer have the 2nd and 5th digits and the metacarpals are fused into cannon bone. The cloven hooves of most artiodactyls (D) represent the evolutionary transition of the 3rd and 4th digits that are now sheathed in keratin. Two exceptions are the padded toes of the camel (E) and the four-toed hippopotamus (F). Senses Artiodactyls are mostly prey animals. As such, they rely on their senses for survival. The eyes are often large and especially adapted to detect movement. Many species may have color vision. Most species have large, mobile ears, and the sense of hearing ranges from good to very good. The olfactory organs are well developed. There is a tactile sense, at least in lips and tongue. Furthermore, there are tactile bristles around the mouth, eyes, and on the checks. Skull and Dentition The skull and teeth vary greatly. In all families except the suidae, it is common to have the front teeth separated from the cheek teeth by a gap or diastema. In the ruminants, the upper incisors are often absent, being replaced by a horny pad. Against this the lower incisors bite when the animal is tearing off vegetation. See family descriptions for dentition. Skulls of four families of Artiodactyla In each of the four families illustrated, the skulls structure has undergone adaptation to provide a survival advantage for the species affected. The elongated, pointed head plates of the peccary allow it to burrow and furrow for food, keeping its sharp incisors and canines in a frontal, attack position. The hippo skull, with its flat cranial plates puts the ryes, nose and ears on top of the head; an ideal position for this highly aquatic mammal. The forward thrusting canines and prominent incisors help in browsing and are excellent attack weapons. The skull of a peccary (Tyassu tajacu) The skull of a river hippo (Hippopotamus Skull length 8.75” (2.5 cm). The elongated amphibius) Skull length 23.5” (60 cm). eve-growing snout adaptation for Forward thrusting incisors and canines are burrowing. major weapons. The skull of a male fallow deer (Dama The skull of a male pronghorn (Antilocapra dama); Family Cervidae; Skull length 10.5” american); Skull length 9.5” (24.2 cm). The (26.5 cm). Both antlers are shed yearly. sheath of the horn is shed yearly, but the bony core is permanent. Note the dentition of the Fallow deer and the Pronghorn. The upper incisors have been replaced by a bony plate, an adaptation typical of ruminants. The Fallow deer and the Pronghorn are primarily prey animals relying on speed and agility as survival adaptations. The antlers and horns are their major weapons and their skulls to provide a suitable platform for their growth. ANTLERS AND HORNS Antlers only occur in Cervidae and then only on the males. The tissue resembles true bones of the body in chemistry composition and cellular structure. Beginning as spikes in yearlings, antlers grow larger each year until prime maturity is reached then decline with age often becoming deformed. Note stages of the Red Deer below. Stages in the development of antlers of the red deer through the life of the animal. Antlers increase in size until prime maturity is reached. a b c d e A B Diagramatic section of antler (A) with velvet and (B) without velvet. (a) velvet; (b) epidermis; (c) dermis; (d) antler; (e) abscission line at burr. Antlers begin to grow in the spring. The developing antler bulges out of the velvet, a hairy skin covering called the pedicel; this is the permanent bony platform atop the frontal bone of the skull. During the period of growth, the velvet is tender and can bleed profusely. About August the antlers ossify, the blood supply and nerves of the velvet are reabsorbed, and sensitivity to injury or pain disappears. The velvet is shed in strips as the animal rubs against trees and shrubs. Once the velvet is removed the bare bone and sharp tines are ready to serve as weapons during the autumn rut in combat with other males for possession of the females. Antlers are shed annually at the end of the mating season, early in December. The cement holding the base of the antler to the pedicel is reabsorbed, weakening the connection. The antlers fall off painlessly one at a time as the animal happens to strike something. Horns are found on the heads of most species of Bovidae, Giraffidae and Antilocapridae. Horns are permanent, ever-growing, and do not regenerate if injured. In species where both sexes have horns, the males are generally larger. The Pronghorn horn is the only one that is branched, otherwise shape varies with species. a b a c b c d d a b c Bovid Horn Pronghorn Giraffe Horn a - keratinized epidermis a - keratinized epidermis a - hair covered epidermis b - epidermis c- dermis b - epidermis c- dermis b - dermis d – bone d – bone c - bone The word “horn” is used to name both the inner bony core and the outer sheath. The outer sheath, the primary component of the horn, is made of keratin, a tough, fibrous protein that forms the outer layer of the epidermal structures such as, hair, nails, horn, hooves, scales, feathers and claws. The sheath is insensitive to pain because it lacks nerves and blood vessels. Bovid Horns: this group includes antelope, cattle, sheep, goats, etc. have permanent outgrowths that are un-forked and horn covered that are usually present in both sexes. Pronghorns have permanent un-branching outgrowths, with a horny covering that is forked and shed annually. Giraffe Horns: in the Giraffe and Okapi, the bony outgrowths are permanent, though they are short, very simple and always covered with skin. The Okapi horns are less prominent than the giraffe. DIGESTIVE SYSTEM In pigs and peccaries the stomach is two chambered and non-ruminating, in the hippopotamus it is three chambered and non-ruminating, In families Camalidae and Tragulidae (mouse deer) it is three-chambered and ruminating. All other ruminants are four-chambered and ruminating. Ruminants, or “cud chewers”, make up the vast majority of the order, including deer, giraffes, pronghorn and all bovids. Diagram of four-chambered stomach Ruminants crop or graze vegetable food, such as grasses and woody material, in which there is relatively low amounts of nutrients they swallow the food rapidly, with little chewing. The food enters the first stomach or rumen (1), later when the animal is resting safely, without fear of predators, this food, already worked on by bacteria, is regurgitated into the mouth, where it is re- chewed and further mixed with salivary juices (the “cud”).